POLSKA AKADEMIA NAUK • KOMITET NAUK GEOLOGICZNYCH acta geologica
PAlIIsTWOWE WYDAWNICTWO NAUKOWE • WARSZAWA polonica
Vol . .27, No. 2 Warszawa 1977
WACLAW BALUK & ANDRZEJ RADWANSKl
Organic communities and facies development of the Kbrytnica basin (Middle Miocene; Holy Cross Mountains,· Central Poland)
ABSTRACT: The Korytnica basin that developed during the Middle Miocene (Badenian) transgression onto the southern slopes of the Holy Cross Mountains, Central Poland, comprises a sedimentary sequence composed of the world-famous, much fossiliferous Korytnica Clays, and of overlying marly sands and red-algal (lithothamnian) limestones. The paleogeographic setting and the structure of the basin, being a part of the drowned valley', are discussed with a special attention to the littoral zone of the basin. This zone was fE'lttured with rubbles and abrasion surfaces, all of which are densely bored by diverse rock-borers (sponges, poly chaetes,. pelecypods, acrothoracican cirripedes). Special littoral facies are exempli fied by the oyster shellbed, and sandy clay with brackish gastropods, the latter of which is interpreted as confined to coastal swamps of the mangrove type. The main facies of the basin, the Korytnica Clays, is discussed in regard with its sedimentary conditions, taphonomy of the fossils and their burial conditions. The most important organic communities and their vertical succession are presented, and a conclusion on the seagrass vegetation in the basin is drawn. Discussed are also some biological relationships between the particular species, and results of the activity of hermit crabs upon alive mollusks. The decline of .sedimentation in the basin manifested with the extreme shallow marine facies that correspOnd to a complete filling of the basin with sediments up to sea level. The climatic conditions that prevailed in the Korytnica i;!asin are characterized as very close to those typical of present-day tropical and/or subtropical zones. Finally, the Indo-Pacific elements within the Korytnica asseglblages are indicated, and a discussion on the connections of the Miocene 'lea of Europe with the lndo-Pacific province is presented.
INTRODUCTION
The aim of the present paper is to show the recent results of investigations on the diversified fossils and their asemblages from the wO!l'ld-famous Middle Miocene (Badenian) Korytnica Clays. These famous 86 WACl.,AW BAl.,UK & ANDRZEJ .RADWANSKI
clays developed within the Korytnica basin on the southern slopes of the Holy Cross Mountains in Central Poland, and the sedimentary sequence of that basin comprised also other facies, all of which are much. fossiliferou~. The fossils from all these fades are commonly regarded as the Korytrtica' fossils, and' under such a name ar,e known in many European collections. 'l1he own research undertaken by the authors in the sixties resulted '''"'in the recognition of general facies conditions prevailing in the basin (Radwailslki 1964, 1969), and of the littoral ic!h.noaoe-noses (RadwaD.s!ki 1970, 1977b); :in the monographing of a part of the mo~lu~ being the most common fossils in the clays, viz. chitons (Baruk 1971), scaphopods (Baluk 1972), and archaeogastropods and some caeoogastropods (Baluk 1975); as well as in the special 'studies on such ' unique fosstils as the creusioid barnacles domiciled in corals (Baluk & Radwanski 1967), and ' the bivalved saooglossan gastropods (Baluk & Jakubowski 1968). More over, during the collecting work, many new groups have been recognized and delivered to the specialists from various countries.' The results of the completed studies on some groups are induded into this issue of ACTA GEOLOGICA POLONICA (Vo!. 27, No. 2; Warsaw 1977). It comprises. the reports on calcareoUs nannoplankton and its stratigraphical importance (Martini 1977), on some large-sized foraminifers (Walkiewicz 1977a), bra.chiopods (Barczyk & Popiel -Barczyk 1977}, free-living bryozoans (Bahik & Radwanski 1977), cuttle fish (Baruk 1977), holothurians (Walkiewicz 1977b), echinodds (Mq- czyiiska 1977), and some fish (Schultz 1977). Moreover, presented aTe . reports on large pinnid pelecypods (Jakubol.yski 1W7), and burrows attributable to the ghost crabs (Radwanski 1977a), both coming from the deposits ' overlying the clays, and finally, a larger paper on the synecology of macrobenthic assemblages from the clays (Hoffman 1977} fills up this isSue. Some other collected groups of the invertebrate fossils are either under investigation (smaJI foraminifeT'S, c()Il'als,tu bular po[yc!haet-es, bryurzoans, cirripedes, crabs, pelecypodls, artimbids, fish oruliths), or ready for the research (os tra codes; ophiuroids, starfi'ldl). If these sltudlies ' a:r'e successful in tJhe forthcoming fu~e, the next . issue on the Korytnica fossils will be kindly offered to the readers.
PALEOGEOGRAPHIC SETTING .OF THE BASIN
The Korytnica basin of Middle Miocene (Badenian) age is a small; c 5 sq km te:rminal part of a larger bay, the Korytnica Bay (cf. Text .,.figs 1-2), Which developed during the. Middle Miocen:~ transgression on the southern slopes of the Holy Cross Mountains in Central Poland .. This. transgression wasoonfined to the area .of the Fore-Ca:rpathian ORGANIC COMMUNITIES AND FACIES DEVELOPMENT 87
Depression (cf. Text-fig. 1B), the development of which had controlled the marine invasion from the Vienna Basin and the Mediterranean (cf. Text-fig. lA).
The Fore-CarpathianDepression, situated at the northern margin of the Carpathiaris (cf. Text-fig. lA-B), is bordered to the north by the Central Polish Uplands (Polish Jura, Cracow and Miechow Uplands, Holy Cross Mts, Lublin Upland - cf. Text-fig. 1B). In a geotectonic -sense, these uplands form the circum -Carpathian belt which was elevated as an iSOstatic response to Carpathian fold ing and formation of the Carpathian foredeep. The .evolution of this foredeep, viz. the Fore-Carpathian Depression, took place in a relatively short time in the Middle Miocene, when the Badenian marine invasion entered this and incorpo rated it into a system of circum- and intra-Alpino-Carpathian basins, commonly called the Paratethys basins (d ~n Text-fig. lA), as they originated from the Tethys Ocean of Mesozoic-Paleogene time. Towards the east, the Middle Miocene (Badenian) transgression reached the western Ukraine. The latter region was very close to the Euxinian Basin (eo in Text-fig. lA) which belonged to the Euxino-Caspian province characterized by more or less lagoonal development with weak connections to the Paratethys basins. Another separate system of basins existed at that time in western Europe where a few Atlantic gulfs (b in Text-fig. lA) encroached upon the continent, and the North Sea Basin developed wider than now {a in Text-fig. lA) and having a temporary at least connection to the Atlantic Ocean through the English Channel (cf. Martini 19'74). On the other hand however, the North Sea Basin had evidently no connection with the Paratethys basins of which the Fore -Carpathian is the northernmost (see B in Text-fig. lA); although unjustified opinion on the existence of such a connection was once expressed by von Linstow (1922) and commonly repeated by subsequent authors (Kauts~y 1925a, b; Sorgen frei 1940, p. 88, and 128; 1958, p. 9; cf. discussion in Friedberg 1930; Gignoux 1955, pp. 592-500 and footnote on p. 603). The triangular shape of northern limits of the Fore-Carpathian Depression (cf. Text-fig. lA-B) results from deep fractures in the Earth crust. The eastern limb, along the Lublin Upland, parallels the SW margins of the Fenno-Sarmatian Shield, being in this part the Ukrainian Platform, and it also parallels the axis of the Danish-Polish Trough (cf. Kutek & Glazek 1972). The western limb of this triangle supposedly· reflects another fracture, more or less perpendicular to the former, both of them being established during evolution of the Danish-Polish Trough, and renewed during development of the Carpathian foredeep. During the Middle Miocene (Badenian) transgression both these limbs were controlling the general extent of the sea which entered the slopes of the Central Polish Uplands along the valleys. The terrestrial erosion . took here place in Paleogene after the Laramide folding of the Danish-Polish Trough and. during successive uplift of the area (cf. Kutek & Glazek 1972). Vadic erosion of surface waters was then accompanied by chemical corrosion of carbonate belts 'Which readily underwent strong karstification in tropical or subtropical climate of the Eocene (cf. Glazek, Dqbrowski & Gradziflski 1972). The valley network was transformed during the transgression into a diversified system of bays with differentiated shoreline, the extent of which is recognizable in the present-day morphology by occurrences of various littoral structures. This is the best de monstrated along the southern slo~s of the Holy Cross Mountains, especially in their western part where the Korytnica basin is located (cf. Text-fig. IB---C). · 88 WACl..AW llAl..UK & ANDRZE.J RADWANSKI
A
'+ ~NIEwrCE 'sOw
5 4 NIZIN~ ~Clolr'~~ .~ " " Oy~ M LESlOWA .
• *JA UPA0.<>v ""sowlEe -PIOT"OWICE ~ A., KORYTNICA,.ARSY rJv~ _o~ F~ IftDSll2DW1CEC-111 sw;:~ ~ Ii.
_&DRI O.._. ~~~8AN1i~~ L ____, 5km~ MY' ~ -- - ~ CHHIWfIJ( . Fig. 1. Paleogeographic setting of the Korytnka basin A - Marine basins in the Middle Miocene of Europe: a North Sea Basip, b Atlantic gulfs I(Brit~any, Touraine and Anjou Basin, Aquitanian Basin, Lisbon Basin), c Western Mediterranean Basin (= Tethys Basin), d Paratethys basins, e Euxinian Basin; rectangled is the area enlarged in Text-fig. 1B B - Extent of ~ the Middle Miocene (Bapenian) sea in the Fore-Carpathian Depression (stippled); rectangled is the area enlarged in Text-fig. 1C C - Middle Miocene (Badenian) shoreline and extent of the bays on the southern slopes of the Holy CrQss Mountains; preserved localities of littoral structures are marked with black spots along the shoreline; asterisked is the Korytnica basin (cf. Text-fig. 2,) situatea in the terminal part of the Korytnica Bay (cf. RadwatiskiD.OO9, Fig. 25; 1970, Fig. 1) Within the inland areas distinguished are the occurrence zones of: 1 Cambrian (including locally Ordovician and Silurian), 2 Devtmian, 3 Triassic, 4 .Jurassic, 5 Cretaceous; marked with haevy dashes are the ridges in morphology that separate particular bays
The valley network was here. established mostly along the strike of the Laramide folds (cf. Text-fig. ic; and Kutek & Glazek 11:l172), built up primarily of the Devonian and Mesozoic carbonate rocks. As result a Dalmatian-type of the shoreline developed in which extensive bays along previous valleys are the typical feature, and these are separated by more or less pronounced rocky ridges (cf. Text-fig. 1C). The mechanical abrasion along the shore produced here such diverse littoral structures as cliffs, cliff rubble, and abrasion surfaces or platforms, all of them having been densely populated by usually gregarious rock-borers (cf. Radwatiski '11964, 1i!169, 19'70, 1974). Within the so-developed bays, the marine Middle Miocene (Badenian) sedi mentation mostly consists of various limestones and marls, the origin of which resulted. from profuse red-algal vegetation. As most of these bays were extensive ACTA GEOLOGICA POLONICA, VOL. 27 W. BALUK & A. RADWANSKI, FIG. 2
Fig. 2. Paleoenvironmental sketch of the Korytnica basin Indicated are: marine area of the Korytnica basin ,during the Middle Miocene , (Badenian) transgression (blank) and present-day outcrops of the Korytnica Clays (stippled); preserved fragments of littoral structures (circled); land or island areas along the seashore (hachured) ASterisked is the summit of Mt. Lysa; marked with black triangle is the summit of Mt. Grodzisko (cf. Text-fig. 3) Numbered are some more important localities discussed in the text: 1 - North-western slopes of Mt. Grodzisko: littoral rubble with borings domiciled by secondary dwellers (Sphenia, Striarca, Creptduta - see PI. 1, Figs 1..,...5); , clay facies ,with brackish gastropods. This is the area of conjectured mangrove swamps (cf. Text-fig. 5), discussed La. by Radwanski (1969. 1974) and Hoffman (19'i'7) , 2 Northern slopes of Mt. Lysa: littoral rUbble covered by oyster shellbed (lumachelles) containing diverse fossils (cf. Text-fig. ,5) described by previous authors 3 The only' natural exposure of the Korytnica Clays, situated on the northern slope of the hill capped by the village, Karsy; this is the locality often named by previous authors as Karsy ' , 4 - Sand pit at Choment6w where the deposits overiying the Korytnica Clays are exposed: these are marly sands and red-algal (lithothamian) limestortes, both containing diverse fossils (cf. Radwanski 1969, Text-fig. '33; 1970, Text-fig. 4; 1977a, b; Jakubowski 1977) Zp - Biotope of the acrothoracican barnacles, Zapfetta pattei Saint-Seine, within the , littoral rubble and fragmentary abrasion surfa,ce (cf. PI. 1;,' 'Fig. 6) ACTA GEOLOGICA POLONICA, VOL. 27 W . BALUK & A. RADWANSKI, FIG. 3
A Mt.Grod zisko
c Chomentow ridge Mt . Wsa
Fig. 3. P resent-day topography along the Middle Miocene ,(Badenian) shoreline of the Korytnica basin (cf. Radwanski 1969, PI. 33; Baluk 197:5, Text-fig. 6) A - View from the village Korytnica (see Text-fig. 2) to localities 1 a nd 2 of the littoral structures (littoral rubble marked with black spots); indicated is also t he present-day extent of the gas tropod-b earing clays (gc ) B - View at the pass between Mt. Lysa a nd Mt. Grodzisko (cf. T ex t-fig. 2); at the background visible is the area of loca lity 1; arrowed a r e fr agm e nts of a brasion surface and littoral rubble; indicated is a lso t he present-clay c:<:e n t of the gastropod-bearing clays (gc) C - View from locality 1 to M t. Lysa (cf. Text-fig. 2); locality 2 is arrowed, an d areas of the gastropod-bearing clays (gc) exposed over the cropland are indica ted; in the back ground visible is the Chomentow ridge bounding the Korytnica b asin from the north (cf. T ext-fig. IC and 2) ORGANIC COMMUNITIES AND FACIES DEVELOPMENT but shallow, and their bottoms were situated within the range of the photic zone, diverse species of the red alga Lithothamnium could thrive here and supply detrital material to various local depoSits. Of these, the most common are pure or ma.rly lithothamnian limestones, and lithothamnian marls composed of algal nodules in the marly matrix. The only exception of such sedimentation type · was the Korytnica Bay which was the deepest of all the discussed bays. Both clastic and lithothamnian deposits were laid down in the open part of the bay, whilst at its terminal, westernmost part, 'behind a submerged ridge a restricted basin had formed. This is the Koryt nica basin I(cf. Text-figs 1C and 2) in which clay material, presumably supplied by a stream or a river from the weathered residues of the Jurassic (Kimmeridgian) clays and marls, was deposited through almost the whole history of the basin: the world--famous Korytnica Clays that contain the extreme wealth of various fossils then originated. Sedimentation of marly sands and lithothamnian limestones appeared in the basin by the decline of · its' history and it completed the sequence of the so fossiliferous strata. Contrary to the region of bays, the eastern part of the Holy Cross Mountains was at that time featured mostly by sandy sedimentation. The clastic material was then supplied from the old Paleozoic (primarily Cambrian) clastics, and sandy shores could widely develop (cf. Radwanski 1970, J.973, 1974).
Within such paleogeographic situation, the Korytnica basin provdded a unique framework to the appearance of very specific fades conditions that resulted both in u~common sediment types, and in the wealth of diversified orgamc remains they · contain.
STRUCTURE OF THE BASIN AND ITS SEDIMENTARY SEQUENCE
The Korytnica basin developed in the part of the ' bay wh!ichwas bordered from 1he north by a ridge built up of massive ... Oxfordian limestones, and from the south, by another ridge built up of LCYwerl IMiddle Kimmeridgian limestones and clays covered peD€COtrdant1y wli.th AlllianlCenomaman compact sandstones that form the top part of this ridge (cf. Text-:tligs le and 2). To the east, another, pr€Sumably submerged 'l'idge separated the basin from the open part of the bay. All these ridges were cut by straits which joined the basin with the other parts . of the bay, and with the open sea to the south, which allowed exchange of water. In result, normal marine conditions prevailed in the basin, and except of some restTficted areas, no oligohaIine oondl tions could persist. The Middle Miocene (Badenian) sedimentary sequence of the basin (cf. Text-fig. 4) begins with local accumulations of the brown-coal de-:. posits, not exposed at the sw-face. The main part of the sequence consists of the Korytnica Clays, the thickness of whiic:h depends on local dendve lations of the substrate, and reaches 40 to 60 m. Overlyjng are sandy marls and lithothanmian limestones, the total thickne&s of which is about 10 m. The presented figures ·of the deposit thicknesses are estima- 90 WACl..AW BAl..UK & ANDRZEJ RADWA~SKI
tive as, it is also ' apparent that a lateral facies variation took place durling sedimentation of the members ov,eTlying the clays (cf. Radwan sk'i. 1969). The Middle Miocene (Badenian) sequence lies in the basin flat (et. Text-fig. 4)" evidencing no tectonic movements except of an isostatic upheaval of the whole basin. Local defoTIllations of some compact layers of marly sands are ascribed to uneven oompa'Ction. The ,present surface that results from Late Tertia,ry und Quaternary erosion, incises various parts of the clays and removes them from the more elevated substrate hummocks making them exposed (cf. Mt. Lysa in' Text-figs 2 and 4-5).
Within the Korytnica Clays there are no larger , exposures, except one at the village Karsy (cf. locality 3 in Text-fi,g. 2). All the paleontological material from the clays is obtained from trenches and local diggings, and from collecting the fossils over the cropland (cf. Text-fig. 3C), especially after spring or fall plowing, the same as after wery heavier rainfall. In last years it appeared that the ablation and resulting concentration of fossils in the soil does not keep up with the acquisitiveness of the collectors.
The westernmosi part of thre basin stretches with a narrow embayment as far as the village Wierzbica (cf. T;€xt-fig. 1C) where the Korytnica Clays have been preserved in some spots and' delivered the same fossils as those :firom the baslin (cf. Kowalewski 1930; Radwanski 1967, 1969).
STRATIGRAPHY
The sedimentary sequence of the Korytnaca baSlm was fm-merly regarded as belonging to the lower part of ,the Polish Tortonian which was cO'I'related exactly with the, Vienna Tortonian (cf. Radwanski 1969, 1973, 1974 a, b). Within the stratigrapbic column of the Polish T,ortonum, there wrere distinguished two substages, the lower of which (Opolian) comprlised a full sedimentary cycle ranging from the transgressive deposits through the final evaporation and formation of gypsum deposits. In such a subdivision, the Korytnica sequence was placed in the lower part of the cycle, i.e. in the Lower Opolian (cf. Radwanski 1969, 1974a).
When new stratigraphic division of the Vienna Tortonian was established, no evident correlations were presented, although it was generally believed that the Polish Tortonian should correspond to the new established Badenian stage of the Vienna Basin.
The fit'st account on the reliable stratigraphic correlations is now presented by Martini (1977) who recognized hi the calcareous nanno plankton from both .the Korytnlica Clays and overlying marly sands, the presence of standard nannoplankton zones NN5 (Sphenolithus hetero- ORGANIC COMMUNITIES AND FACIES DEVELOPMENT 91 morphus Zone) and NN6 (Discoaster exilis Zone). In result, MartiIlli (1977, Text-fig. 3) demonstrates that the Korytrcica sequence is cOrrela table with the upper part of the classical section of c1aysat Sooss, and with the lower part of the WalbersdOTf section in the Vienna Basdn. When taklmg into account the current biostratigraphdc subdivision of the Central Paratethys Mtooene (Steininger, Ragl & Martini. 1976), it is evidenced by Maa-tini (1977) fi:1at the KorytnicasequeIlIce is the equi valent to the middle part of the Vienna Badenlian, anid to the upper part of the type Langhian. in Italy; it cannot be li:n any case correlated with the type Tortonian of Piechnont in which much younger nannoplankton zones (NN9 - NNll) are present (cf. aLS9 Martini 1975; Steininger, Ragl & Martini 1976).
THE LITTORAL ZONE OF THE BASIN
The littoral zone of the Korytnica basin is shaped mostly by littoral rubbles encountered on the slopes of . the shore, alpng which small areas of abrasion surfaces developed locally (cf. Text-figs 2-3). Usu ally, abrasion surface was formed in parts of the shore where more compact and resistant limestones were exposed, whereas 'fue layered limestones were trahsformed into the littoral rubble (cf. Text-figs 3 I;ind5). All these littoral. structures are densely boTed by diverse rock -borers, among which the most important aTe (cf. Radwanski 1964, 1965a, 1969, 1970): sponges CUona celata Grant and C. vastifica Han cock; polycllaetes Potamilla reniformis (0. F. Miiller) , Polydora ciliata· (JoIhnston) and P. hoplura (Claparede); pelecypods Gastrochaena dubia (Pennant), Aspidopholas rugosa (Brocchi), Jouannetia semicaudata (des MouHn:s) , and Lithophaga lithophaga (Linnaeus); 8JrlId acrothoracican barnacles ZapfeUa pattei Saint-Seine. All these rock-borers are very common, locally even gregarious, in all the sites of littoral struc~es / ' (cf. PI. 1), except of the acrothoracican barnacles ZapfeUa pattei Saint- Seine whic'h appear in greater quantities only in one place (marked Zp in Text-fig. 2; cf. PI. 1, Fig. 6).
Of these rock-borers a special attention is to be paid to pelecypods whose shells,or internal moulds, are sometimes preserved within the borings. They allow to determine both the boring species, and to ascribe the empty borings to particular species as the morphology of the borings strictly corresponds to that of the shells (cf. Radwanski 1965a; 1969, 1970, 19'77b). The most. common material within the ichnocoenose, the empty borings of the pelecypods, may , therefore be well ascribed to normal body-fossil species. All the remaining taxa of the borings fall into the category of trace fossils.
When comparing the ,composition of the littoral ichnocoenoses of the Korytnica basin with those of the open-sea parts of the Korytnica 92 - WACl..AW BAl..UK & ANDRZEJ RADWA~SKI
bay, and of other regions of the Ho~y Cross shores, it is evident (cf. Radwanski 1969, 1970) that some rock-borers (mostly echinoids) are missing ftn this protected part ·of the bay, and others (e.g., polychaetes Polydora . hoplura, and pelecypods Lithophaga lithophaga, especially their larger forms) -are much less frequent here. These differences re sulted from hydrodynamic conditions within the particular biotopes and from biological requirements of these rock-borers (cf. Radwanski 1969, 1970).
INHABITANTS OF EMPTY B9RINGS
Many of the empty pelecypod bodngs within the littoral rubbles, _especially those at the locality 1 (cf. Text-figs 2-3 and 5) were domi ciled by various secondary dwellers whose shells remained preserved (cf. PI. 1,Figs 1-5), the same as of primary rock-borers (cf. PI. 1, Fig. 3). The squatters of empty borings are mostly (cf. Text-fig. 5; and . Rad'wanski 1969, 1970, 1974 a, b) pelecypods Sphenia anatina (Basoorotj. and Striarca lactea (LinnaeulS), the latter of which has a similar mode of life in present-day environments in the Adriatic and Black Sea (cf. StaTmiihlner 1963a, Nevesskaya 1965). Lessfreq~ent are the gastro pods, the slipper limpets Crepidula crepidula (Linnaeus), Which more commonly occur as secondary dwellers of any empty shells within the Korytnica Clays (cf. PI. 2, Fig. 15; and Radwanski 1969, 1974b). All these squatters of empty borings adopted their shape to that of the bo:dngs regardless their outlinecorrespond1ng to diverse rock ·boring pelecypods. In result, the shape of the squatters is much variable and deviated from the "atandard" forms of particular species. Some times, it was also dependant upon the space left by the original rock -borer whose shell remained preserved in the boring during the life
KORYTNICA CHOMENTOW
Fig. 4. Idealized section through the Korytnica basin (not to vertical scale; cf. Text-fig. 2) to show the general sequence of the Middle Miocene (Badenian) deposits 1 brown-coal deposits, 2 KORYTNICA CLAYS, 3 marly sands, 4 red-algal (lithothamnian). limestones ACTA G E OLOO1CA POl.ONiOA. vor_ ...
'(jJ;;, C C",-",ity II Ij (j) "l' 1JJ L ,,.. ,.•• L!!GANll: •• pebbl••. """.~ ""~ oy Id""lized SOOre9CO.p e to ...'l of a squatter (cf. PI. 1, F'iig. 3; Sphenia anatina adopted to the shell of Jouannetia semicaudata preserved in its boring). OYSTER SHELLBED The oyster lumachelles, composed of various shell detritus, mostly of the oyster Ostrea frondosa de Serres and other pelecypods associated with gastropods, and contained in marly matrix, are a local littoral fades (cf. Friedberg 1928, 1931; Radwanski 1964, 1969). This facies is best developed on the slopes of Mt. Lysa which made up an !island . during sedimentation in the Korytnica basin (cf. Text-figs .3-5). On the northern slopes of Mt. Lysa, there appears the only exposure of this facies which covers a littoral rubble, as it is visible in a small abandoned quarry (localiJty 2 in Text-:Hg. 2; cf. also Text-fig. 3A and C) . . The oyster lumachelles form a kind of the oyster shellbed (os in Text-fig. 6) which was accumulated simultaneously with the deposition of the Korytnica Clays further off the island shore. The oyster rriaterial vanishes at the distance of c 30-50 m off this shore, and the deposit gradually becomes the gastropod-bearing Korytnica Clays (cf. Text -fig. 3A). AJlthough nOlW:the oyster sheUbed wraps up the whole island of Mt. Lysa(cf. Text-fig. 6), the oyster banks during sedimentation were certainly developed locally, mostiy amidst elevations of the substrate which, being more exposed, became habitats fur the rock-borers (cf, Text-fig. 5). In other parts of the shore slope the seagrass carpets then also occurred. (cf. Text-fig. 6). A rich community of the oyster' banks comprises also (cf. Text-fig. 5) diverse . oorals" mostly Dendrophyllia taurinensis Milne-Edwal'ds .& Haime, BalanophyUia vari!Zns Reuss, and others (cf. Dembiilska-R6zkow ska 1932, Kiihn 1963), as well as cirriped'es of the genera Scalpellum Leach, Balanus . MANGROVE COAST Another littoral facies of a local development is the clay with greater admixture of quartz sand, and containing commonly some gastropod species which are either rare or missing in the main area of the Koryt nica Clays.. This sandy clay is exposed only on the north-western slopes of Mt. Grodzisko (locality 1 in Text-figs 1-2), and it covers a littoral rubbl'€' with the above.;.discussed secondary dwellers of empty pelecypod borings. . 94 WACl.AW BAl.UK & ANDRZEJ RADWAN"SKI The gastropod species such as Terebralia bidentata (Grateloup), Pire neUa tabulata (Homes), Cerithium aff. zelebori HOI"lnes, Neritina picta Ferussac, and Melanopsis aquensis Gralleloup, are of typical brackiish cha!racter. Their occurrence here (cf. Text-fig. 5) may be explained by special paleogeographi'C oonditions that, controlled a local development of the restrliiCted environment. It was a part of the shore of the ridge (cf. Text-figs 1C and 2) that wa's facing the island of Mt. Lysa (cf. Text-figs 2-3). The water agitation and circulation were here certainly weaker, which explains . the preservation of some shells of rock~boring pelecypods, and of secondary dwellers within the emptied borings (cf. RadwaIlski 1969, 1974). On the other hand, the morphology of the discussed ridge featured wiith broad denivelations that paralleled marly horizons 'within the Jurassk substrate, produced a notchy shoreline with dlifferentmted facies ooll1ditioll1s. The re.rrestrial material, both clayey and sandy,. was easier accumulal1Jed here, and it blurred and oovered' qukkly the rocky substrate. It was suggested (Radwanski 1974) that under such conditions the coastal swamps could develop, and they were overgrown by mangrove-type thickets (cf. Text-fig. 5). In such a spot, a prOltected , environment might have been established amidst' th~ quaggy ponds and the maze of pirop roots (cf. Plaziat 1970, Taylor 1971, Braithwaite & al. 1973), and the braokish conditions prevailed even if the fre:sh-water . supplieS' frOllIl the diSICUssedridge have nolt beien inuneniS!e (cf. alSo Plaziat 1975 a, b). The above interpreta1JiJOn on the occurrence of mangrove swamps (cf. also Hoffrnan 1977) is partly based on that presented by Braithwaite & al. (1973) for some members of the Quaternary sequenJCe of th~Aldabra Atoll, Indian Ocean, where the presence of Terebralia and other PLATE 1 1-5 - Squatters of emptyborings in pebbles making up littoral rubble at Korytnica. (locality 1 in Text-fig. 2; cf. also Text-fig. 5) I Gastropod Crepl.dula crepidula (Linnaeus) adapted to the boring of Gastrochaena; at right, pelecypod Striarca lactea (Linnaeus)- squats another boring (marked L); X 1.5 2 Gastropod Crepidula crepidula (Linnaeus) adapted to the boring of Ltthophaga provided with a crest corresponding to the hinge margin of the boring shell (cf. Radwaiiski 1969, PI. 35. Fg. 8); X 1.5 ' 3 Gastropod Crepidula crepidula \~mnaeus) adapted to another boring of Ltthophaga; above, pelecypod Sphenia anatina (Basterot) squats (marked S) a small b'oring of Jouannetia semicaudata (des Moulins)' whose preserved shell is also visible (marked J) 4 Pelecypod Sphenia anatina (Basterot) adapted to' the boring of Gastrochaena (cf. Rad- waiiski 1969, PI. 35, Fig. 9); nat. size , 5 Pebble with pelecypods Sphenia anatina (Basterot) domiciled in diverse borings, mostly of Lithophaga; the largest shell is visible projecting out of the partly' damaged boring (ct.Radwaiiski 1989, PI. 35, Fig. 10); ,nat. size 6 -'- Swarm of borings of the acrothoracican barnacles, ZapfeZZa pattei Saint-Seine, within the abrasion surface bored by pelecypods (cf. Radwanski 19.69, PI. :trr, Fig. 2); Korytnica (locality marked Zp in· Text-fig. 2); nat. size ACTA GEOLOGICA P OL ONICA, VOL. 27 W . BALUK & A . RADWANSKI, PLo 1 ACTA GEQLOGICA POLdNICA, VOL. 27 W. BALUK & A. HADWAJii'SKI,- PLo 2 Typical gastropods of the Korytnica Clays 1 - AnciUa g!andiformis (Lamarck); 2 - Murex friedbergi Cossmann & Peyrot; 3 - Tudic!a rusticu!a (Basterot); 4 - Sigaretus striatus de Serres; 5 - Natica mmepunctata Lamarck; 6 - Cypraea !anciae Brusina; 7 - TurrtteUa badensis Sacco; 8 - Genota va!eriae (H. Hoernes & Auinger); 9 - Conus ponderosus Brocchi; 10 - Aporrhais a!atus (Eichwald); 11 - PyrUla geometra Borson; 12 - Fusus virgineus sensu Htirnes, and sensu H. Hoernes & Auinger; 13 - Triton nOdiferum Lamarck; 14 - Terebra acum inata Borson; 15 - HaneUa papi![osa Pus ch, the shell contains a group of the slipper limpet Crepidu!a crepidu!a (Linnaeus): large female and two p ygm y males, X 1.5 ; 16 - Clavatula !aevigata (Eichwald) . All figures in nat. size, except of Fig. 15 ORGANIC· COMMUNIT~ES AND FACIES DEVELOPMENT' 95 cerithids is regarded as indicative of restricted circulation, and the presence of mangroves (cf. Braithwaite & a1. 1973, p. 311). The compa rable gastropod communities of the present-day mangrove swam.ps are also reported from Madagascar, Maur;.tius, Mozambique and western Mrica (cf. review and references in Plaziat 1970). THE KORYTNICA CLAYS During sedimentation in the Middle Miocene (Badenian) sea, the Korytnica Clays were' cOvering almost the whole area of the basin (cf. Text-figs 2 and 4), although at present they are exposed on a rather strongly limited aiI"ea (stJippled in Text-fig. 2). In the I'Iemaindng paxt of the basin, at the tops of morphologic elevations exposed now at the surface there are the members overlying the clays (maTly sands and red-algal limestones.; cf. Text-fig. 4), whereas in depressions ' used by streams of the present-day outflow, the Pleisrocene gla.cral outwash sands, and Recent black muddy soil make up the cover of the clays (cf. Text-fig. 2: blank are all the areas where the clays are 'not exposed a t the surface). The Korytnica Clays are of light yellow color, and they are well recognizable at the surface, especially when a deeper plowing was done. After rainfall, such places become whitish due to a great content of s.hell detritus, mostly of the small pelecypod CorbuZa (AZoides) gibba (Olivi). Larger gastropod shells are easily perceptible also in the soil over the whole cropland that features vast areas of the basin (cf. Text-fig. 3A-C: areas marked gc). ·· The Korytruica Clays are soft, plastic, and no bedding or any other physical sediimentary structures Me discernible, except of local lami natiO!llcau'Sled by sihell detritus. Due to the lack of any oontrast between successive portions of the clays, neither burrows nor bioturbations are recognizable, although plenty· of various endobenthic animals, La. gastropods and pelecypods (cf. Hoffman 1977), and some hol,othurians (cf. Walkiewicz 1977b) lived in the basin during depoSition of the clays. To begin with presenting the history of investigation of the Korytnica Clays, it should be noted that recently the occurrence of the fossils reworked from the . substrate rocks has been indicated. It concerns the microfossils, both calcareous 'nannoplankton reworked from the Jurassic and Cretaceous deposits (Martini 19,77), and some holothurian sclerites reworked from the Upper Cretaceous (Wal kiewicz 197'7b). If a delivery of the Jurassic fossils with the fine clastic material is quite obvious, as discussed above (chapter on the structure of the basin), the supply of the Cretaceous material has to prove a transport from the open sea areas, presumably through the strait in the place where at present the village Korytnica is situated (cf. Text-fig. 1C and 2). 96 WACl.AW BAl.UK & ANDRZEJ RADWANSKI HISTORY OF INVESTIGATION The history of inveetigation of the Korytnica Clays has long been the history of recognition of their fossils (cf. Radwanski 1969, Baluk 1975), mostly gastropods,- as this very group makes the most common and outstanding componeDlt of the orgaruc woirld within 1:\he clays. These gastropod-bearing clays (cf. Text-fig. 3 and PI. 2) have often been called the Pleurotoma Clays, as the pleurotomids, mostly of the genus Cla vatula if not the commoneSt (cf. PIs 2-5), are ,certamJ.y O!llle of the most striking gastropods of the Korytnica Clays. The locality of Kory1Jnica was discovered presumably somewhere about the twenties of iihe last century (cf. Kowalewski 1930, Baluk 1975), but the f4"st description of !its fossils appeared some twenty years later (Pusch 1837). . An international career 'of Korytnica began in the forties .of the last century, when Sir R. 1. Murchison on his famous way to Russia stopped in Poland specially to make a trip to this locality. His brief account otn the locality and collected gastropodS (Murdhioon 1845, pp. 292-293) completed that presented formerly by Pusch, and it was the first description of th~regional situation of the clays. The Korytnica gastropodS were later included by Eichwald (1853), Hornes (1856) and Hoernes & Auinger (1879) into their classical mono graphs of the Mibcene cmalacofalina. The sucMollusca miocaenica Poloniae", published in seven parts (gastropods and scaphopods 1911-1928; pelecypods 1934-1936; guide to the collection 1938; for references to smaller contributions see Ba.luk 1975; revision of the genus Conus - see Hall 1964). Both Dr. K. Ko walewskl and Pl'ofessor W. Frledberg were the first who paid attention to other fossils than mollusks, either pUblishing own contributions (cf. Friedberg 1924, 1930), or delivering the fossils to specialists in the country and abroad (e.g., polychaetes - Dembiilska 1923; COTals - Dembinsika-R6zkowska 1932; fish otoliths - Chaine & Duvergier 1928).· The scien1ific activity of Professor W. Friedberg and of Dr. K. Kowa lewski closed the period precediJl1g that of the .recent investigators. REVIEW OF THE ANIMAL GROUPS The Korytnica Clays contain the fossils representing almost all the systematic groups of the and.m.al kingdom (of vertebrates only fish are present) whose skeletal parts are recognizable in the ancient deposits. The state of their reoogni1fion and chances for further research are as follows. ORGANIC COMMUNITIES AND FACIES DEVELOPMENT 97 Foraminifers The abundant foraminifers have not hitherto been investigated systematically, except of the miliolids (Luczkowska 1974), and of some contributions on selected groups or genera , (cf. Baluk 19<75; Walkiewicz 1975, 1977a). Radiolarians Infrequent radiolarians have not hitherto been studied systematically. Sponges The boring sponges are only rec'ognized (cf. Radwanski \119064, 1969, 1970; and this paper), as well as the presence of some others domiciled by cirripedes Acasta and/or by gastropods Tenagodus is inferred (cf. Baluk & Radwanski '1967, Baluk 1975; and this paper). Corals The corals, both solitary and colonial, were recorded by most of the previous authors (cf. Baluk 19175). They were included into the monograph of the Polish Miocene corals by Dembinska-R6zkowska (1932). The genera Flabellum and Acanthocyathus were afterwards discussed by Kojumdgieva "1960), and/or by Kiihn (HI63) while the creusioid inhabitants in the species ·Tarbellastraea reussiana (Milne-Edwards & Haime) were studied by the authors (Baluk & Radwanski 1967; cf. also this paper). Recently, the corals are subjected to investigation by Dr. E. Roniewicz, Institute of Paleobiology, Polish Academy of Sciences, Warsaw. Polychaetes The polychaetes are represented by boring forms {cf. Radwanski 1964, 1,9119, 1970; and this paper), as well as by various tubular, either free-living (Ditrupa). or epizoic (Spirorbis, Serpula) forms (cf. Dembinska 1923, Baluk 1975; and this paper). Bryozoans The bryozoans are recognized very imperfectly (cf. Baluk 1,975). The ecologi cally importaqj; genus Cupuladria, reported from here previously (Cook 1965, p. 199~, is the subject of a separate paper by the authors (Baluk & Radwanski 1977). Special ~udies on the Korytnica bryozoans a;re recently kept by Professor J. Malecki, School of Mining and Metallurgy, Cracow. Brachiopods The previous reports on the brachiopods were very scant (cf. Friedberg 1930, Kowalewski 19'30, Baluk 1975). This group is now. monographed by Barczyk & Popiel-Barczyk (197'r). Ostracodes The ostracodes halVe not hitherto .been systematically studied '(cf. Baluk 19.75), 98 WACf.AW BAf.UK & ANDRZEJ RADWA~SKI Crabs The predatory activity of the hermit crabs upon the alive '.mollusks is a common feature of the Korytnica fossils (Radwanski 19169, L977b; cf. also this paper). A rich material of the claws" the only preserved parts of the crabs within the clays, is now investigated by Dr. R. Forster, University of Muni~h. Cirripedes A great asemblage of 'Various cirripedes, containing representatives of common acorn barnacles and such genera as Scalpellum, Conchoderma, Verruca, Chtha malus, Acasta, Creusia, some of which have not hitherto been known from the Miocene (Conchoderma, Chthamalus), is the 'subject of the authors' own investi gations. The first result, the paleontological and ecologic analysis of the Creusia species has been included into the monograph of the Miocene cirripedes domiciled in corals (Baluk & Radwanski 1'967). Further investigations are in progress (cf. Radwanski 1969, 1·974a, b; Baluk 197.5; and this paper). The etching activity of the genus Verruca is presented separately (Radwanski 1'9'77b; cf. also Baluk 1975, and this paper). The trace fossil species, Zapfella pattei Saint-Seine, attributed to the acrothoracican barnacles was recognized within the littoral ichnocoenoses of the basin (Radwatiski 1964, 1969, 1970; cf. also Baluk 1\9175, and this paper). ChltoDS The assemblage of chitons, including fifteen species, and being therefore the richest in the European Miocene has been recently monographed by Baluk (19'71; cf. also this paper). Sca.phopods The assemblage of scaphopods, including thirteen species, and being one of the richest in· the European Miocene, has also been monographed by. Baluk (1972). Gastropods The most important component of the gastropod-bearing clays (cf. history of inv~stigation) comprises an assemblage of about 800 species, some of which are new. This becomes the subject of a new monographing by Baluk; the first volume has just recently appeared (Baluk 19176), and the others are in preparation. Separately described by Baluk & Jakubowski (1968) was the bivalved sacoglossan species of the genus Berthelinia; whose bearing on the facies conditions within the basin have been often reminded (Radwatiski 1969, 1974a, b, 1975; cf. also this paper). The predation by the muricids and naticids upon other mollusks, and their mutual competition in the finding and selecting of prey were studied by Hoffman & al. '(1974), whereas mortality patterns of some species were described by Hoffman (1976b). ' Pelecypods The other important component of the clays <;omprises an assemblage of about 200 species, much richer than it was formerly ' recognized {cf. Kowalewski ORGANIC COMMUNITIES AND FACIES DEVELOPMENT 99 1930; Friedberg 1931, 1932, 1934-1936, 1938). This assemblage is now investigated by Dr. G. Jakubowski, Museum of the Earth, Polish Academy of Sciences, Warsaw; he has recently included six Korytnica species into his monograph on the ontoge~ic development of .some pelecypods from the Polish Mioeene (Jaku~ bowski 1'972), and contributed on unique large pinnids from the mar1y sands overlying the clays '(Jakubowski 1977). The rock-boring species, the same as secondary dwellers · of empty · borings were studied by Radwatiski {1'964, 1965a, lQ.69, 19'70, 1977b; cf. also this paper). Mortality patterns of some species have recently been presented by Hoffman (il976a). Cephalopods Of the cephalopods, only the presence of the cuttlefish has recently been recognized by Baluk (197'5), and subjected to paleontological description of a new species (Baluk 1977). Crlnolds The presence of the fl,"ee~living comatulid crinoids of the genera Antedon and Discometra has recently been recognized by Baluk (1975) who obtained them from siftings and now is finishing their description. Holothurlans A very rich assemblage of the holothurian sclerites is recently monographed by Walkiewicz (197.7b) who recognized eleven new species apart from some twenty common species. The presence of some other holothurians is inferred from the, synecology of organic communities, and from some biologic . relationships (see below). Ophiuroids and starfish Infrequent ossicles of both ophiuroids and starfish are collected Echinoids The echinoids were very little known formerly (cf. Kowalewski 1930, Ba luk 197'5), but now they became the subject of their monographing by Mq czytiska (1977). F:ish The fish otolitbs are rather common material in the clays, and they have long been the subject of interest {Friedberg 1924, Chaine & Duvergier 1928, Smigielska 1966); their newvmonograph is prepated by Dr. T. 8migielska, School of Mining and Metallurgy, Cracow. The tooth material, both of the · elasmobranchs and teleosteans, previously reported very accidentally (cf. Kowalewski 1930, Baluk 1975), and moreover .some bone · material of the teleosteans· are- now . monographed . by Schultz (1977). 100 WACl..AW BAl..UK & A~R:ZEJ RADWAN"SKI ANIMAL COMMUNITIES During sedimentatiOIli of particular horizons of the Korytnica Clays the bottom conditions, except of the littoral zone (see above), were almost unifOrnl all over the basin (cf. Radwanski 1969, Hoffrnan 1977). The same may be said when regarding the type of the bottom sediment that was laid down in the vertical succesSion of the clay member (see above). The fossil content in the vertical succession is however much differenJtiated, and it reflects the Changes in organic (mostly mollusk dominated) communities which lived during the gradual filling of the basin by the clay deposits (cf. Radwaiiski 1969, 1974a, b). The. vertical succession of organic communitieS in any part of the clay sequence corresponds well to that established along the shore slope when the basin was formed (see idealized shorescape in Text-fig. 5, and succession of the communities in Text-fig. 6). The distribution of the fauna along the shore slope resulted then ev1Jdently from bathy metric requiremell1ts of particular genera or species, as follows. The community of the deepest part of the shore slope (Community l' in Text-fig. 5) is characterized by a solitary coral Flabellum (cf. Dembiilska-R6z kowska 1002, Kojumdgieva 1000, Kiihn 19163), large tusk shell Dentalium (AntaZis) badense Partsch (cf. Baluk 19'72), and gastropods Turritella (cf. Baluk H)'75). This community is comparable to those typical at present of the depths ranging between 40 and 60 meters (cf. Radwailski 1969). The community of the middle part of the shore slope {Community 11 in Text-fig. ·5) is characterized by the abundance of diverse gastropods, the most typical of which are various Clavatula, Murex, Ancilla, Conus, Cypraea, Tudicla, Strombus {schematically drawn in Text-fig. 5), as well as Turritella, Cassis, Triton, Fusus, Ch en opus, various Natiea and Nassa (cf. PI. 2). Less common are Ranella, Pyrula, Genota, Terebra, an,d Sigaretus (cf. PI. 2), as well as the slipper limpet Crepidula crepiduZa :(Linnaeus) which inhabited empty shells of any other gastropods, and is often preserved in groups' composed of a larger female and pygmy males perching on her (cf. PI. 2, Fig. 15, and PI. 3, Fig. 6). Associated are colonial corals, mostly Tarbellastraea 'reussiana {Milne-Edwards & Haime), and various pelecypods, whereas numerous incisions in shells show the remarkable role played by the hermit crabs {cf. PIs 10-12; discussion below). The rock-borers, the same as those occurring in the littoral zone (see below), appear in the shells within the discussed biotope I(ef. PIs 3-5). There also appear the fragments of large-sized mollusks, mostly gastropoos (cf. PIs '1-8) of the genera Triton, Murex, Conus, VolutiZithes, Cypraea, Cassis, Rostellaria, Strombus, Xenophora, and Galeodes, whose presence makes a separate problem (see below). The whole Community 11 is comparable to those typiCal at present of the depths ranging between 20 and 40 meters '(cf. Radwailski 196~). T~e cOQlmunity of the highest part of the shore slope (Community III in Text-fig. 5) is generally very similar to the preceding one, although many of the gastropods acquire larger size and thicker shells. Moreover, some new animals appear, the most attractive of which is the bivalved gastropod Berthelinia brow sing on the kelp (the only locality of this gastropod in the European Miocene; cf. PI. 9, Figs 1-3, and Baluk & Jakubowski 196,8), various chitons with Craspedochiton and worm-shaped Cryptopla~ I(cf. PI. 9, Figs 6-7; and Baiuk 1971), ORGANIC COMMUNITIES AND F ACIES DEVELOPMENT 101, cirripedes Creusia domiciled in corals (cf. PI. 9, Fig. 8; and Baluk & Radwanski 1967), as 'Well 'as Verruca, ChthamaZus and common acorn barnacles Balanus. The discussed genera live today along rocky coasts or in coral reefs just below low water level (some of them even spreading up into the intertidal zone). The whole Community III is therefore regarded as living at the depths shallowing from some twenty to a few meters (cf. references in Baluk' 1971, Baluk & Jaku- ' bowski 1968, Baluk & Radwanski 11967, Radwanski 19169). / The whole sequence of the Korytruca Clays, as shown by the above characteristics of the organic communities and their life condit ions, should consequently be interpreted as formed in the suocessively shallowing basin. Due to gradual storage of the clay deposit, the depth of the basfu. was successively decreasing, and the organic communities, previously dlstributed laterally at various depth ranges of the shore ,slope (cf. Text-fig. 5) could spread over the whole basin to repeat vertically their former lateral array (cf. Text-fig. 6). The original depth of the basin was, of that value to which the pre-Miocene valley (cf. Text-fig. le) was drowned durfug the transgression, whereas the final portion of the clays was deposited at the extreme shallow depths, but s:tlill within the submerged mnes (cf. Text-fig. 6 and 'See belOlW: ohapter on seagrass vegetation). SEAGRASS VEGETATION The analysis of the whole animal aSlSemblage of the Korytruica Clays shows that the -presence of the Sieagrass meadows must be taken into account when conBiderting the synecologic relationships. 'TIhis' is apparent both :from the compOsition of particular communities which are well comparable to those ct>nfined to seagrasses in present-day environments (Hoffman 1977), espedfullyof the Indian Ocean (cf. Taylor 1971), and from the indis,tinct etchings on the mollusk shells, which are regaIl.'ded as counterparts of those left by seagrass roots in the present-day habitats (cf. Ho:ffman 1977, Cort:treU 1977, Radwanski 1977b). Additional argumentation is delivered by a common presence, in some habitats at least, of the animals which in present-day environments live primarily within the seagrass spots. This is true for such groups~ or particular genera and species, as: cuttlefish (cf. Baluk 1977; and Starmiihlner 1963b); free-living comatulid crinoids of the Antedon type (cf. Strenger 1963a); some holothurians (cf. Walkiewicz 1977b; and Stren ger 1963b); echinoids Echinocyamus which have recently been recorded from the -clays (Baluk 1975, Mqczytiska 1977) and which are m06tly confined to such habitats (cf. Szorenyi 1953, Strenger 1963c). Sirruilar conclusion is also drawn from the presence of the foraminifer species 2 102 WACf.AW BAf.UK & ANDRZEJ RADWAJ.Il"SKI Amphistegina lessonii d'Orbigny which lived tprough the almost whole time of deposition of the clays (cf. Walkiewicz 1975; and Bandy 1964, Rosenkrand's Larsen 1976). c kelp seagrass sea level kelp seagrass I~ Fig. 6. Succession of the organic communities (1, 11, 111) during the formation of 'the Korytnica Clays (c:!. Text-fig. 5) A - Time after deposition of the lowermost part of the Korytnica Clays (characterized by the community I) and a contemporaneous part of the oyster shellbed (os); communites 11 and III spread over higher parts of the submerged slope, the same as the seagrass and kelp B - Ti.me after deposition of the middle part of the Korytnica Clays (characterized by the community 11) and a cqntemporaneous part \ of the oyster shellbed (os); community 111 spreadS over higher parts of the submerged slope; the seagrass meadows cover the bottom almost entirely C - Time during depositon of the uppermost part of the Korytnica Clays (characterized by the community 111) and a contemporaneous part of the oyster shellbed (os); the seagrass . meadoWs still cover the· bottom almost entirely, and locally stretch out of the sea level · ORGANIC COMMUNITIES AND FACIES DEVELOPMENT 103 The seagrass commun.ities durJng . deposition of the lowest part of the Korytnica Clays were thriving mostly along the submerged slopes of the seashore where the photic requirements were sufficient for their vegetation (cf. Text-fig. 6A). The seashore slopes were then occasionally covered by seagrasses, whereas in the remaining ar'eas either rocky substrate was stretching out of the sediment surface, or the oyster shellbedwas formed. Later, when the basin was successively shallowing, the seagrass communities were spreading throughout the basin (cf. Texf-fig. 6B), and such a situation lasted until the uppermost part of the clays was deposited. At that time the seagrass meadows could reach sea level in some places (cf. Text-fig. 6C), in a: way similar to ,that at the tops of mud banks on the Flor;i!da offshores (cf. Ginsburg & Lo- wenstam 1958). The same situation is also recognizable for the 1:i.me of depositiO!I1 of the marly sands that overlie the KorytniCa Clays (cf. Rad wanski 1977a). FOSSILS OF THE KORYTNICA CLAYS In this chapter presented are such new data on the distribution, ecology, taphonomy, and state of preservation of the fossils from the Korytnica Clays, which have a bearing upon general cOnclusions on the Life and preservation conditions within the Korytnica basin. DISTRIDUTION All the fossils' within the Korytnica Clays are clistr'ibuted very Ir-..· _ ·- .. -· .. ·- · --•• ----, ,ununiformly. As it is apparent in trenches, the fossils are often ooncen trated in streaks or thin lenses in which the frequency of parti,cular species much varie~. La'l'ger gastropods are to be foun!dsometim~ in groups on small areas, whereas in ' ot1?-er places they occur very sporadically and are dispersed singly in the clay. The abundance of gastropods within all the successive hotTizons of the Korytnica Clays is certainly remarkable, as they are often re pr€Sented by numerous specimens of ' a few species of the same genus (e.g. Clavatula, Natica, Nassa); Either these ' species were of different ecological requirements withiln the same biotope, or the shell. accu- . roulation in the deposit came from various biotopes of lateral and/or vertical succession. It seems that generally the necrocoenOS€s correspond to life habitats of the gastropods, as no sigruificant post-mortem sorting or transportation is indica:t:ed (cf. Hoffman 1977). Nevertheless, rut is difficult to recog:nire whicih species lived quite contemporaneously, as the particular n€CTotopes conJtain the species not only epibenthic, but also (e.g. Natica) endobenthic. On the. other hand, a local segregation 104 WACl.AW BAl.UK & ANDRZEJ RADWANSKI and transportation of Shells became an important factor, especially outside the seagrass meadows. Finally, when regarding the shell distribution, such biological ag,ents. of transportation should be taken into account as the activity of scavengers, and of the hermit craoo using the shells as their homes. TApHONOMY The burial conditions during sedimentation of the Korytnica Clays were much differentiated and, if no significant transportation of the shells is kept in mind, they should be ascribed mostly to a variable sedimentation ratio. Most of the shells contained in the Korytnica Clays are fresh, and these were supposedly buried just after the death of the host. There are however oommonly met various shells more or less corroded, or .strongly damaged due to the organic activity. The corrosion is expressed by a frosty surface, usually of one side that was stretching over the sea floor. Some sh~lls are bored by rock-borers in the same way, similarly as those encrus'ted by the epizoans (see below). It is therefore apparent that such shells were partly buried 1x:> a definite level, presumably quickly; and afterwards they were stretching naked with their upper parts out of the sea floor for a longer span of time. Such shells became then the sedimentary traps for s'maU-sized fossils which concentrated here in comparison with the clay adjacent to the entrap ping shell, and which met here exceptional conditions for their pre servation. This is well demonstrated for juvenile pelecypods (cf. Jaku bowski 1972) and brachiopods (ef. BaTczyk & Popiel-Barczyk 1977), chitons (cf. Baluk 1971), some tiny gastropods (cf. Baluk 1975), and for free-living bryazoans (cf. Baluk & Radwanski 1977). On the other hand, the preservation of some gastropod shells inhabited by the slipper limpet Crepidula erepidula (Linnaeus), the female of which often carrii~s one or a few pygmy males on her shell PLATE 3 Activity of boring sponges in the mollusk shells of the Korytriica Clays 1-5 - Borings of CZiona vastifica Hancock in the gastropod shells: 1 - Lyria taurinia (Bonelli), 2---'i - AnciZZa gZandiformis (Lamarck), 4 - Ocenebra erinacea . (Linnaeus), 5 - Purp,ura exiZis Partsch; all X 1.5 6-10 - Borings of CZiona ceZata Grant in the gastropod shells or their fragments: 6 - ClavatuZa .asperulata I(Lamarck) - the shell is partly cut-off to show the slipper limpet Crepidula crepidula (Linnaeus) settled inside (arrowed): large female with a pygmy male; 7--8 - Ancilla glandiformis {Lamarck), 9 - fragment of Xenophora deshayesi (Michelotti), 10 - fragment of Triton nodiferum Lamarck; all X 1.5 11-13 - Clionid borings in the pelecypod shells or their fragments: 1'1 - fragment of Spondylus crassicosta Lamarck, and 12 - fragment of Chlamys latissima nodosiforms (de ·Serres), both bored by Cliona cei(J.ta Grant; 13 Chama gryphoides garmeZla de Gregorio bored by Cliona vastifica Hancock; all in nat. size . ACTA °GEOLOGICA POLONICA, VOL. 27 ': ' W: -BALUk &' ° A. RAbWA~SKt , PLo ~ ACTA GEOLOGICA POLOij'ICA, YOL. 2'1 W. BALUK & A. RADWAN'SKI, PLo 4 ,' . ORGANIC COMMUNITIES AND FACIES DEVELOPMENT 105 (cf. Fig. 15 in PI. 2, and Fig. 6 in PI. 3), evideIliCes no transportation, a.nd a rapid burial. It may therefore be concluded that the shells contained in the Korytnica Clays. were lying on the sea floor for a various. span of time: some were buried very quickly, while some others underwent a gradual destruction (see below). The latter ones could certainly then became the- components of such necrocoenoses to wffiich new shells were temporarily delivered. Recently it is suggested (Baluk 1977) that of the nectk cuttlefish, preserved in the clays were the skeletal parts of only these individuals . which, during · their seasonal migration to the Korytnica basin (see below), had beco,me the prey of the predators. It may be also noted that the burial conditions dUl'ling sedimentat- 'ion of the Korytni'ca Clays w&e favorable for preservation of color pattern in some fossils. This is well recognizable in some gastropods (cf. Baluk 1975), the bivalved Berthelinia including (cf. PI. 9, Fig. 3a; and Baluk & Jakubowski 1968), as well as in the creuSibid barnacles (cf. PI. 9, Fig. 8; and Baluk & Rad'wailski 1967). ENDO- , AND EPIZOANS The shells of the fossilS contained in the· Korytnica Clays often became the habitat of various endo- and epizoans, the ac1ftvity of w!hich was either p['onouIlJCed on one side of the shell (as ind'icatled above), or on its whole (cf. PIs 3-6). 'Dhe endbzoans aTe represented generally by the same rock-borers whioh are present iin littoral rubbles and abrasion· surfaces (cf. Text -fig. 5), except of the acrothoracian barnacles (cf. Radwailski 1969, 1970). Their damage in the mollusk shells is either . local, or very total and resulting in their fragmentatlon into pieces. PLATE 4 Activity of boring polychaetes in the mollusk shells of the Kortynica basin 1-5 - Borings of PoZydora ciliata (Johnston) in the gastropod shells from the Korytnica Clays: 1~ - CoZumbeZZa curta (Dtijardin) - the boringsexposed to show their position paralleling the adapical channel of the last whorl; 3 - Natica' redempta Michelotti - vertical borings in the callus; 4 - AnciZZa gZandi formis (Lamarck) - borings extended around the aperture; 5 ~ another AnciZZa gZandiformis - shallow borinl1s on the outer surface of the shell; all X 1.6 6--11 - Borings of PoZydora hopZura (Claparede) in the gastropod shells from the Korytnica Clays: 6 ---: CZavatuZa Zaevigata (Eichwald) - openings of numerous borings are visible; 7 - CZavatuZa asperuZata (Lamarck), and 8 - C. Zaevigata - the borings exposed io show their position within the columella and the last whorl; 9 - Cerithium vuZgatum miospinosum Sac co - a group of borings exposed; 10 - AnciZZa gZandijormis, and 1i1 - fragment of Euthria puschi (Andrzejowski) - in .both the borings are exposed; all X US 12 - Numerous borings of PoZydora hopZura The boring sponges, Cliona celata Grant and C. vastifica Hancock, lived in gastropod and pelecypod shells (PI. 3, Figs 1-13), resulting often in their total damage (cf. PI. 3, Figs 3 and 7-12). Some small fragments,. strongly bored by sponges (cf. PI. 3, Figs 9-12) certainly come from the shells heavily destroyed which have therefore easlIY been broken into pieces. The boring polychaetes, Polydora ciliata (Johnston) and P. hoplura (ClaparMe), settled mostly in the gastropod shells (PI. 4, Figs I-H), while in the pelecypod shells this second species is common, but it occurs mostly in the marly sands overlying the clays (PI. 4, Fig. 12). These two polychaetes were boring at various depth and at various angle to the shell surface (cf. PI. 4, Figs 5-10), some of the borings having been sligthly incised into the shell (PI. 4, Fig. 5). Larger bo-rings, those of P. hoplura, are often situated along. the thickef>t part of the shell, e.g. along the columella (PI. 4, Fig. Bp. In smaller borings, those of P. ciliata, it is ·not so common, although in the shells having the callus, this part is bored primarily (PI. 4, Fig. 3). Some borings. of P. ciliata are situated along the aperture of the shell I(PI. 4, Fig. 4) which may evidence their origin during the gastropod life. This latter possibility should mostly be taken into account in the species Columbella curta (Dujardin), the shells of which contain often only one boring of P. ciliata, but it continues along the adapical channel (PI. 4, Figs 1-2). Of the pelecypods boring in the littoral zone of the basin, only Gastrochaena dubia (Pennant) is present in the mollusk shells of the Korytnica Clays (cf. PI. 3, Figs 1--,9,). Sporadically, borings of Petricola sp. may also be found in the clays (Radwatiski 19690. The gastrochaenid borings are well attributable to the· species. indicated above, G. dubia (Pennant), as their shells are often preserved in the borings (cf. PI. 5, Fig. 8b). Many of these borings were only partly embedded in the shell,and their upper parts were covered by ;in agglutinaceous tube; when the latter have been removed, the "half borings" appeared from beneath (cf. PI. 5, FigS" 2 and 4-5). Agglutinaceous is also the siphonal collar that outlined the siphons stretching out of the shell surface (cf. PI. 5, Figs 1~2), the same as out the rock surface in the littoral zone (cf. PI. 5,Fig. 3). The mode of formation of the agglutip.aceous parts of the tube that may complete the partial borings is the same as in present-day gastrochaenids, the species G. dubio, (Pennant) including i(cf. references in Radwanski 11969; also Warme J.9'15, . Fig. 11.25). The frequency of the gastrochaenid borings on one shell varies from a single PLATE 5 Activity of boring pelecypods, Gastrochaena dubia (Pennant), at the rocky seashore, and in the mollusk shells of the Korytnica Clays 1-2 - Borings of Gastrochaena. dubia (Pennant) provided with the agglutinaceous siphonal collar. that stretches out of the shells of Ancilla glandiformis (Lamarck); in 2, upper part of the boring wall has not been preserved (this is the "half boring"); X 1.5 3 - Agglutinaceous siphonal collar stretching out of the rocky surface; an epizoic coral is visible nearby (cf. Radwanski J.009, PI. 34, Fig. 3; 1970, PI. 3b); X 2 4 - Numerous "half-borings" in Conus sp.; X Hi 5 - "Half-boring" oriented tangentially in Clavatula asperulata (Lamarck); X 1.5 6-'-7 - "Half-borings" oriented vertically in: 6 - Cypraea lanciae Brusina, 7 - . Purpura exilis Partsch; X 1.15 . 8a - Terebra fuscata (Brocchi) with a few borings, in one of. which (arrowed) was preserved the shell of Gastrochaena dubia (Pennant) presented in 8b; both . X 1~ 9 - Vertically oriented borings in Glycymeris pilosa deshayesi (Mayer); nat. size • ACTA GEOLOGICA P OLONICA, VOL. 27 W . BALUK & A. RADWANSKI, PLo 5 ACTA GEOLOGIC A POLONICA, VOL. 27 W. BALUK & A. RADWA~SKI , PLo 6 ORGANIC COMMl1NITIES AND F ACIES DEVELOPMENT 107 occurrence (cf. PI, '5, Fig. 1) to several specimens (cf. PI. 5, Figs 4 and 6-8) and larger groups of the borings, especially within the iarger shells of pelecypods (cf. PI, 5, Fig. 9). A general conclusion as to the endozoans in the mollusk shells of the Karytnica Clays is ' that they occur in all the horizons of the clays, and therefore the depth of the baSin was convenient for settlement of the littoral rock-bore.rs through the w!hole time of clay dieposlitidn. The rock-borers which inhabited the shore slope during the transgression (cf. Text-fig. 5) were always, able to settle in any hard objects, i.e. the mollusk shells when they appeared in the areas of clay sedimentation. l Within the mollusk shells there is no preference to any rock-bore.rs, . and almost all the recognized species have been tinfested to some extent.• An average value of the shells infested in particular species is however low, and it may be estimated as about 3-5%. The activity of the epizoans within the mollusk shells , of the Korytnica Clays, the oyster shellbed including, is recognizable by the presence of etchings (cf. Radwanskrl. 1977b) produced by. the cirripedes of the genus Verruca when their shells were attached (PI. 6, Fig. 3a-3b; cf. Radwanski 1974b, 1977b). The other, deep etchings which may also be regarded as borings (PI. 6, Fig. 4), belong to the bryozoa'll ' genus Spathipora, the systemati:c ~tudies on which are now kept by the authors. Indistinct etchings left by seagrass roots (see above) complete the list of the hitherto recognized taxa. Some . epizoans are preserved with thefur hard parts, as it is exemplified by tubes of Spirorbis sp.t (PI. \6, Figs 1-2) and other 1 Specific determination of these spirorbids is not obvious at the moment. Previously reported from the Korytnica Clays was only Spirorbis ' obtectus Seguenza, listed by Kowalewski {19~O, p. 71 and 1'11) who uncorrectly suggested its biological relationship to the gastropod AncilZa glandiformis Lamarck, on the shells of which it was found. This spirorbid species was inCluded by Rovereto into the synonymy of Spirorbis (Laeospira) umbiliciformis (Munster), but such ' an assignation has not been accepted by Schmidt (1:95'5). The investigated specimens are partly comparable to those of the latter species, and presented by Schmidt (1955; PI. 8, Fig. 32). PLATE 6 Activity of encrusting and etching invertebrates in the ' mollusk . shells of the Korytnica Clays 1 - Polychaetes Spirorbis sp. and a bryozoan encrusting Lyria taurinia (Bonelli); encrusted is the apertural side of the shell bored by clionid sponges (its opposite side being presented in PI. 3, Fig. 1); X 1.5· . 2 - Group of polychaetes Spirorbis sp. encrusting· the inner side of the shell of Tudicla rusticula (Basterot); X , 5 . 3a - Shell of the gastropod Lemintina arenaria (Linnaeus) from the oyster shellbed ,(the specimen presented by Baluk 197~, PI. 14, Fig. 14) encrusted by an oyster (oy) end etched by verrucid barnacles, Verruca sp. the most etched part (arrowed) is magnified in 3b; nat. size 3b - Part of the shell presented in the preceding photo, to show the pit-shaped etchings produced by Verruca sp., (cf. Radwatiski 1977b, PI. 8c'); X 5 4 - Deep etchings prqduced by bryozoans Spathipora sp. in the shell . of Ancilla glandiformis (Lamarck); X 5 108 WACl..AW BAl..UK & ANDRZEJ RADWA~SKI polychaetes, as well as by very uncommon bryozoan colonies (PI. 6,. Fig. 1) and corals, the latter being also known from the littoral zone (cf. PI. 5, Fig. 3). Within the oyster shellbed with Ostrea frondosa de Serres, many shells are encrusted by this very oyster (cf. PI. 6, Fig. 3a). OCCURRENCE AND PRESERVATION OF LARGE MOLLUSKS The presence of very large-sized moliusks, mostly gastropods, is a very characteTis1Jic feature of the middle and upper part of the Korytruica Clays (cf. Ra'dwaIlski 1969, p. 103; 1974a, p. 112). In these members of tlhe clays., i.e. wi1llrin, the commUJn!i:tLes Il and III (cf. Text -fig. 5) such gastropod shells are enough commonly found, but, all are preserved i'D; fra~ents (cf. Pls 7-8). Some of 1lhem. are strongly damaged by rock-borers (e.g., PI. 7, Fig. 7b; PI. 8, FU.gs 1 anid 3a), the same as those occurring both ill 1ihe ' littoral tzanle and in "normal" sized shells (cf. PIs 1, and 3--4), but others are quite fresh. Most of the fragmented ,large gastropods bear remarkedly thick shells, and their so sev'ere damage :is therefore puzzling. Of 1ihe hitherto recognized gastropod species, all here discussed are much larger than the undamaged specimens of these species from the Korytnica Clays, and some of them Me even larger than the largest ones from the other European'localities (mostly of the Pairatethys basin - cf. Text-fig. lA), as follows:, Triton nodiferum Lamarck - the colleeted specimens (PI. 7, Fig. 1) belonged to individuals attaining c185-190 mm in their height, being larger than those from the Vienna Basin (cf. Hornes 1856). Similar larger forms are however known from the Pliocene of Italy and Cyprus. A mould of the specimen of comparable size (c 170 mm) was formerly reported also from the marly sands that overlie the Korytnica Clays (Radwanski 1969, p. 106). Cassis miolaevigata Sacco the collected specimen (PI. 7, Fig. 2) was c 65 mm high, being therefore of the size comparable to that of the Vienna specimens (cf. Homes 1856). Volutilithes haueri (Hornes) - the collected specimen tPI. 7, Fig. 3) was c 70-75 mm high, being much larger than those from the Vienna Basin (cf. Hornes 1856, Hoernes & Auinger 18719), Hungary, (cf. Strausz 1966), and Bulgaria (cf. Kojumdgieva 1960). J Cassis cypraeiformis Bprson - the collected specimen .(PI. 7, Fig. 4) was c 80 mm high, being also rpuch larger than those from the- Vienna Basin (cf. Homes 1856, Hoemes & Auinger 1879) and Hungary (cf., Strausz 1966). Rostellaria dentata Grateloup - the collected specimens (PI. 7, Fig. 5 a, b) belonged to indi'Viduals attaining c 180 mm in their height, being therefore of the size comparable to that only known in specimens from Baden in the Vienna Basin (cf. Homes 1856). Strombus ,coronatus sensu R. Hoernes & Auinger - the collected specimens (PI. 7, Fig. 6) belonged to individuals attaining c 100 mm in their height, being of the size almost identical ~with that in specimens from the Vienna Basin (cf. Homes 1856), and from Lapugy in Transylvania, (cf. Hoernes & Auinger 1879). ACTA GE OL OGICA P OL ON ICA, VOL. 27 W . BALUK & A. RADWA~SKI , . PL . · ~ \. Fragments of the shells of large gastropods from the . Korytnica Clays (cf. explanation to PI. 7; the r everse side of this fold-out); nat. size 1- 2 - Diverse Conus sp., 3 - GaZeodes cornutus (Agassiz), 4 - Murex austriacus Tournouer, 5 - X enophora deshayesi (Michelotti) ACTA GEOLOGICA POLONICA, :VOL. ~ 7 W. B l'. LUK & A. RADWAI'<' SKI, PL. 7 Fragm€nts of the shells of lar ge gastropods from the Korytnica Clays; the hypotethical outline of the shell is shown as taken from the comparative material (some fragments dp . not match exactly these outlines, being derived from the specimens of morphological ratios others than that one used to draw the outline); nat. size 1 - Triton nodiferum Lamarck, 2 - Cassis miolaevigata Sacco, 3 - Volutilithes haueri (Hornes), 4 - Cassis cypraeiformis Borson, 5 - RosteHaria dentata Grateloup, 6 - Strombu.s coronatus sensu R. Hoernes & Auinger, 7 - Cypraea sp. ORGANIC' COMMUNITIES Al\"'D FACIES DEVELOPMENT 109 Cypraea Sp. - the collected specimen (PI. 7, Fig. 7) was c 70 mm high, which howe;ver does not reach the size of various large cowries from t~e Vienna Basin (cf. Hornes 1856, Hoernes & Auinger 1B7f1). Diverse Conus sp. - the collected specimens (PI. 8, Figs 1-2) certainly belonged to different species; a fragment of the spire (PI. 8, Fig. 1) is not eVidently determinable, whereas a fragment of the last whorl (PI. a, Fig. 2) presumably belonged to Conus daciae Hoernes &'Auinger, being of the size (c 90 mm) compar able to that reported by Hoernes & Auinger (1879) from 1f;he Vienna Basin and Lapugy in Transylvania. GaZeodes cornutus (Agassiz) - the collected specimens I(PI. 8, Fig. 3 a, b) belonged to individualsattainingc 1-60 mm in their height, being therefore the largest bulbous, thick-shelled, and thus the "heaviest" gastropods of the Korytnica basin, although they were much smaller than those from Niederkreuzstatten in the Vienna Basin (205 mm high; cf. Hornes 1856), and from the world-famous collection site of this species at Varpalota in Hungary (c 220 mm high; cf. Strausz 19 ' 66)~ Murex . austriacus Tournouer - ,the collected specimens {PI. 6, Fig. 4) belonged to individuals attaining c 100 mm in their height, being therefore a little larger than those from Enzesfeld in the Vienna Basin {cf. Hoernes & Auinger 187!9~. Xenophora deshayesi (Michelotti) - .the illustrated specimen (PI. 8, Fig. 5), and many other fragments of similar size, belonged to indilviduals attaining even c 100 mm in their width (e.g., the specimen in PI. 8, Fig. 5, the outline of which is drawn smaller than it should be taken from· that fragment - camp. the lowest part of the photo). Such specimens were evidently' wider thap the widest ones from the Vienna Basin (cf. Hornes 1856); although they were of the size of those from Hungary (cf. Strausz l.!l1(6). The two of the above listed species, Volutilithes haueri (Homes) and' Galeodes cornutus (Agassiz), aTe hitherto known from the Koryt nica Clays only from such damaged fragments. It was previously suggested (Radwanskl 1969, p. 103; 1974a, p. 112) that the discussed large gas1;rlopods presumably lived within the littoral rubble where they underwent damage by grinding between stones in the surf zone, and then the fragmen'ts were swept away the .shore. T1his may explain why the two above-mentioned species are unknown from the cxffshore facies, but it connot clar'ify Why some fresh-looiking fragments are found in areas distan.t to the Shore, where from no evident transport is indicated (cf. Hoffman 1977). It seems therefore that even the species with the thickest Shells lived far away the shore. If so, the only assumed cause of their total damage is the activity of predators, supposedly of that kind as huge crabs, or large vertebrates such as some :fisih or mammals 2. The large-sized pelecypods from the Korytn~ Clays certainly underwent similar damages as the gastropodS' did, although they occur ! In this respect, even the presence of terrestrial or semi-aquatic mammals may also be taken into account, as indicated by a commonly referenced example of the Californian sea otter which, when dived up the shell, floats on his back nearshore and hammers this shell against the stone he carries on his stomach! (cf. Bishop 19'75, p. 265). -, 110 WACl,AW BAl,UK & ANDRZEJ 'RADWAN'SKI very sporadically and never make an essential ',content of thenecro coenose. They are r,epresented by a few species only, such as: Glycy-' mer is pilosa deshayesi {Mayer} whose isolated valves aTe often bo['ed by Gastrochaena gp. (cf. PI. 5, Fig. 9); Cardium hians danubianum Mayer and Meretrix gigas (Lamarck) preserved in small fragments; Chama gryphoides garmella de Gregorio Whose isolated valves are da maged by clionid sponges (cf. PI. 3, Fig. 13), the same as small frag ments (cf. PI. 3, Figs ll-i2) of Spondylus crassicosta Lamarck and . Chlamys latissima nodosiformis (de Serr,es), the latter species of which has not hitherto been reported from the Korytnica Clays. BIOLOGIC :EtELATIONSHIPS Some biologi.cal relationships whkh persisted in the ba;sin during deposition of the Korytnica Clays are well demonstrated by natural associations whi.Ch have been preserved in their life relation. This is shown the best by the creusioid barnacles, Creusia sanctacrucensis Bafuk & Radwa-:h&ki which lived, as commensals, within alive colonies of the corals Tarbellastraea' reussiana (Milne-Edwards & Haime) that occur in the middle and upper part of the clays (cf. PI. 9, Fig. 8; and Baluk & Radwaflski 1967). Other _examples of similar relationships are inferred from the presence of one species, the partI1€r of which remaiI1€d however' unp~eserved. This is well demonstrated e.g. by the cirripedes Acasta whJicih have be:en srtated in 1Jhe oyster sh:ellbed (cf. Baluk & Radlwait.ski 1967, p. 498), and which in p!esent-day environments are exclusively adapted to live embedded in the sponges (cf. Darwin 1854), The same partner, a sponge, is inferred from the presence (cf. PI. 9, Fig. 4) of 'PLATE I} Some ecologically important (commensal or thermophilic) invertebrates from the uppermost part of the Korytnica Clays 1-3 - Bivalved gastropods, Berthelinia kraehi Baluk & Jakubowski; well discernible ..color pattern displayed by radial stripes visible in 3,a; left valves, to show the protoconch - a outer, b inner view (cf. Baluk & Jakubowski 1008, PI. 1); X ~O 4 - Aberrant gastropod Tenagodus (Tenagodus) anguinus mioeaenieus Cossmann & Peyrot living in a sponge (cf. Baluk 1975, PI. 14, Fig. 16); collected in" the uppermost part of the oyster she11bed; X 8 5 - Aberrant gastropod Parastrophiq, radwanskii Baluk (cf. Balukl1975, PI. 14, . Fig. 1); X 16 6-7 - Chitons Cryptoplax weinlandi. Sulc: 6 head 'Valve, 7 intermediate valve (et. Baluk 1971, PI. 6); X 115 8 - Aberrant cirripedes, Creusia sanetaerueensis Baluk & Radwanski, domiciled in corals Tarbellastraea reussiana (Milne-Edwards & Haime); five well preserved, and two damaged crowns are visible, ' all of them having well discernible color pattern displayed by concentric stripes (cf. Baluk & Radwanski 1967, Text-fig. 3 and PIs 1-6); X 5 . ACTA GEOLOGICA POLONICA, VOL. 27 W . BALUK & A. RADWANSKI, PLo 9 ACTA GEOLOGICA POLONICA, VOL. 27 W. BALUK & A. RADWANSKI, PLo 10 Results of activity of the hermit crabs on the shells of alive gastropods Clqvatula laevigata {Eichwald): a - undamaged shell, b - "kitchen midqens" left by the crabs on the sea floor (cf. Radwanski 1977b, PI. lla); nat. size ACTA GEOLOGICA POLONICA, VOL. 27 W. BALUK & A. RADWANSKI, PLo 11 Results of activity . of the hermit crabs on the shells of alive gastropods Tudicla Tusticula (Basterot): a - undamaged shell, b - "kitchen middens" left by the crabs on the sea floor (cf. Radwanski 1977b, PI. lIb); nat. size ACTA GEOLOGICA POLONICA, VOL. 27 W . BALUK & A . RADW ANSKI, PLo 12 ORGANIC COMMUNITIES AND FACIES DEVELOPMENT 111 the gastrowd Tenagodus (Tenagodus) anguinus miocaenicus Cossmann & Peyrot, which supposedly lived within the oyster banks in the same way as in present-day environments (cf. Wenz 1939, p. 680; Baluk 1975; Morton & Miller 1973, 2 in Text-fig. 74). As discussed above, the vegetation of the kelp Caulerpa in the littoral zonle is ooncluded from the presence (cf. PI. 9,Figs 1-3) of the bivalved gastropods Berthelinia krachi Baluk & JakubowskLThe present -day representatives of this genus live only in the habitats populated by this v1ery kelp on which they browse (cf. Keen & Smith 1961, Baluk & Jakubowski 1968). The presence of some more holothurians than those indicated with sclerites by Walkiewicz (197-7b), is recognized by the trophic analysis of some mollusk assemblages (Hoffman 1977) in which the gastropod genus Eulima was certainly adopted to ectoparasitic mode of life on sea cucumbers. Others are evident fro:gl the presence of the fish otoliths Carapus nuntius (Koken) recently recognized by Dr. T. Smigiel ska (pers. information), and Which are attributed to the genus Fierasfer, the present-day repr,esentatives of which live (cf. Zei 1963) in the anus of the holothurians Stichopus, or rarely of Holothuria. The presence of the hermit crabs, inferred from the damages in molluSlk sihells, is discussed in the following chapter, whereas a conside ration on possible predators of the free-living hryozoans is omitted here (cf. Baluk & Radwanski 1977). ACTIVITY OF HERMIT CRABS . The -activity of hennit crabs on alive mollusks from the Koryt nica basin is the best pronounoed within the riddle and upper part of the- Korytnica Clays, i.e. within the communities II and III (cf. Text -fig. 5 and Radwaiiski 1969, 1977b), The most severe damage caused by hermit crabs within these communities is confined to gastropod species provided with long-siphoned shells, such as primarily Clavatula laevigata (E'iohwald) and Tudida rusticula (Basterot) whose shells are often deeply incised by crab claws, and cut mto halves (cf. PIs 10-11).' The reSulting "wra!cks" of the PLATE 12 1 - Results of activity of the hermit crabs on the shells of alive gastropods AncilZa glandiformis (Lamarck) 2-13 - Incisions produced by the hermit crabs, and regenerated by the mollusks (cf. Radwanski 1969, PI. 37; ;197'1b, PI. 12): 2 - scaphopod Dentalium fossiZe raricostatum (Sacco), and the ' gastropods: 3 - Nassa schoenni (R. Hoernes & Auinger), 4-5 - Columbella curta (Dujardin), 6 - Euthria puschi {Andrzejowski), 7 - Turritella badensis Sacco, 8- Sveltia inermis (PUsch), 9-lO and 12 - ~lavatula laevigata (Eichwald), 11 . and 13 - Clavatula asperulata (Lamarck) 112 WACl.AW BAl.UK · & ANnRZEJ RADWANSKI shells are commonly met in this interval of clay sedimentation and, if selected from the associa.ted unattacked shells, they look like the "kitchen middens" left by satisfied crabs on the sea floor (cf. Radwan ski 1977b). Nevertheless, the particular species of the Korytnica gastropods were attacked by hermit crabs to a very variable extent. Of the long -siphoned fo!l'Il1s, the most common two Clavatula species, C. laevigata (Eicihwald) and C. asperulata (Lamarck), became 1lhe prey in very low percentage, whereas rather uncommon Tudicla rusticula (Basterot) was damaged in most of the specimens collected. A low percentage of the damages :is also noted in a common species AnciHa glandiformis (La marck) whose thick shells may however be nipped completely (cf. PI. 12, Fig. 1). When investigating the particular species, succeSSive stages of the crab nippings are recognizable (cf. Radwanski 1969, 1977b). Generally, unsuooessful for the crabs were incisions not deeper than about a half Whorl, and these were regenerated by the hosts that survived. The l'\egenerated shells, both of gastropods and scaphopodS are recogndzable in most of the common spedes (cf. PI. 12, Figs 2-13) 3. If a deeper incision by the crab had reached the mantle, and damaged it, the subsequent regeneration was associated with vaTious anomalies in the shell shape or sculpture (cf. Radwanski 1969, 1977b). The presented results of the hermit crab activity in the Koryt niea basin do not differ from those previously reported both from present-day and Neogene environments (cf. Papp & aL 1947, Boekschoten 1967, RObba & Ostinelli 1975, BiShop 1975), being generally indicative of shallow marine conditions (cf. also Radwanski 1969) . . OTHER PROBLEMS During recent investigation of various fossils from the Korytnica .Clays, an attention has. been paid to some new problems which may certainly be fully recognirzed by further I1esearch. When studying the ecology of the collected sepioids, Baruk (1977) noticed that these animals might have migrated into the shallows of the Korytniea basin for breeding by springtime. A similar sugges:tion was presented by Dr. T. Smigielska (pers. information) for some bathyal . fish whose otolitihs she recognized in the Korytniea Clays. For the two animal groups, namely some regular echinoids (Mq- czyilska 1977) and free-living bryozoans (Baluk & Radwailski 1977) it was stated that the clayey bottom of the Korytnica' basin was thedi- 3 Except for the two above-mentioned species, TudicZa rusticuZa (Basterot) and AnciZZa gZandiformis (Lamarck) which have never been found regenerated. These two species therefore either could not regenerate their shells, or all the crab attacks upon them were successful. - ORGANIC . COMMUNITIES AND FACIES DEVELOPMENT 113 favorable environment, otherwise to that stated for the comparable present-day forms. The latter is demonstrated especially for tlJ.e free -living bry()lZ()an species Cupuladria canariensis (Busk) which is extant, and whose biology in present-day environments is recognized the best (cf. Lagaadj 1963, Cook 1965, Baluk & Rad'wanski 1977). MEMBERS OVERLYING THE CLAYS The members overlying the Korytnica Clays, i.e. marly sands (3 in Text-fig. 4) and red-algal (li1fuothamnian) limestones (4 in Text -fig. 4), both exposed the best in; a sand-pit at the village Chomentow (locality 4 in Text-fig. 2) complete the sedimentary sequence of the Korytnlca basin. Their lithology, fades development and paleontological content lis! now discussed by Radwanski (1977a), and may be summarized as follows. The marly sands that gradually appear at the topmost part of the Koryt~ nica Clays, and make up the transitional beds c 1.5-2 meters thick (cf. Rad~ wanski 1969, Baluk '1975), contain diversified fossils. These are mostly large foraminifers, Heterostegina costata d'Orbigny and single colonies of red~alga Lithothamnium. Typical is an assemblage of pelecypods, containing both small cockles, Cardium paucicostatum Sowerby, and such large~sized species as Panope menardi rudolphii Eichwald, Meretrix islandicoides (Lamarck), M. gigas (Lamarck), Cardium hians danubianum Mayer, Crassostrea gryphoides {Schlotheim), all of them being commonly reported by former authors {cf. Kowalewski 1930, Friedberg , 1930, Radwanski 1969). Recently recognized by Jakubowski (197-7) is a new species of the pinnid genus Atrina, formerly· assigned to the genus Pinna itself (Rad~ wanski 1969, 1970). Associated are lingulid brachiopods {Friedberg 1930, Baluk 1975, Barczyk & Popiel~Barczyk 119177), various echinoids (Baluk 1975, M~czyn~ ska .19'17), mostly of the genus Echinocyamus, starfish Astropecten (cf. KO'Walew~ ski W30, Baluk 19715), and large gastropod Triton nodiferum Lamarck (the latter mentioned above). Large oysters are here usually encrusted by serpulids and acorn barnacles (Radwanski Hlta9). The bottom was, partly at least, stabilized by seagrasses (cf. chapter on se.agrass vegetation; and Radwanski 1977a). Indicative of the bathymetric conditions of that member are the burrows attributable to the ghost crab Ocypode, the presence of which suggests a very shallow :rn.a!Tine environment, just below or even within the intertidal zone (Radwaflsk.i 1977a, b). These burrows, which occur in the upper part of the sands (cf. Radwaflski 19'Z7a, Text-fig. 1)~ evi dence therefore an extreme 'Shallow marine (possibly,' brackish-water influenced; cf. Wallciewicz 1975), or temporarily emerged episode in the history of the Korytnica basin, that is, its almost complete :ffi1ling with tlJ.e sedimentS'o The red-algal (lithothamnian) limestones which overlie the sands, record a I'Ieturn of maTine sedimentation and a rise of sea level to about 5-6 meters (cf. Radwaflski 1977a). The basin was then successively 114 WACf..AW BAf..UK & ANDRZEJ RADWANSKI filled with algal limestones in which embedded are large cobbles and boulders derived from the shore, i.e., from the Chomentow ridge (cf. Text-figs 1C, 2 and 3C), distant c 200-300 meters (cf. Text-fig. 2). The largest bould,ers, attaining over 1 meter in their diameter, that occur in the topmost paTt of the limestones (cf. Radwanski 1977a, Text-fig. 1) indicate a hurricane transport over :sLippery algal meadows when they have carpeted the basin almost up to sea level (Radwanski 1969, 1970, 1977a, b). The boulder-bearing horizon marks the final epiSlOde of the total filling of the Korytnica basin with sediments and completes its marine history. It is presumed (Radwanski 1969; cf. also 1974a, 1977a) that the Korymica basin, certainly together with the shallow bays of the Holy Cross shores (cf. Text-fig. 1C), w.ere then definitely excluded from the marine sedimentation which however still lasted in the deeper, open-sea facing part of the Korytnica' Bay, and in the remaining areas of the Fore-Carpathian . Depression. CLIMATIC CONDITIONS Within the organic oOmmunities of various members of the Koryt nica basin there appear many animals which are indicative of tropical and/or subtropical climatic conditions. This is the best demonstrated by the taxa recognized fu the Korytnica Clays, as follows. The genus Creusia, adapted to living in alive colonial corals, and to which the species C. sanctacrucensis Baluk & Radwatiski belongs (cf. PI. 9, Fig. 8), is distributed at present primarily within the reef corals I(cf. Baluk & Radwatiski 1967, Ross & Newman 1973, Text-fig. 2). The bwalved gastropod genus Berthelinia, which is represented (cf. PI. 9, Figs 1-3) by the species B. krachi Baluk & Jakubowski, lives at present in tro pical and/or subtropical lit~oral zones only, where it browse on the kelp Caulerpa growing mostly inbetween rubbles of strongly agitated waters just below low water level {cf. Keen & Smith 1961, Baluk & Jakubowski 11958). Of the untvalved gastropods, such genera as Terebra, Rostellaria, Architec tonica, and large-sized species · of the genera Cypraea, Conus, Strombus, Triton, Voluta (Volutilithes), an;:l Galeodes are commonly regarded as typical of tropical waters. The same is also true for the genus Parastrophia, repre~ented (cf. PI. 9, Fig. 5) by the species P. radwanskii Baluk; the genus being formerly unknown in the fossil state (cf. Baluk 1975), is confined at present to the tropical zone (cf. Wenz lro~, Baluk 1975). . Of the chitons, represented are the. species of the genera Craspedochiton. and CryptopZax, both living at present in the tropical zone, and the latter of which, being worm-shaped {cf. PI. 9, Figs 6-7), is adapted to living within cre vices in coral reefs and other littoral rocks in the tropics (cf. Ladd 1966, Baluk 19171). Of the foraminifers, instructive is the species Amphistegina lessonii d'Orbigny, the present-day representatives of which are confined to tropical and subtropical waters (cf. Walkiewicz 1975, Rosenkrands Larsen 1976). The same ORGANIC COMMUNITIES AND F ACIES DEVELOPMENT 115 is said about the free-living bryozoan species CupuZadria canariensis (Busk) which at present has similar climatic requirements (cf. Lagaaij 1963, Cook 1965, Cadee 197'5, Baluk & Radwanski 19,77). Tropical and/or subtropical character is also displayed by some holothurian sclerites which are ascrib€d to the present-day holothurian genera of those cli matic zones (cf. WalkieWicz 19!77b). Finally, to the same climatic zones are confined the mangrove swamps (cf. Plaziat 1970), and some mollusk communities, to which the Korytnicaassem blages are compared (cf. Hoffman 1917). In the marly sands, a similar bearing upon the climatic conditions is displayed by the burrOlWS attributable to the ghost ~ab Ocypode which at present is distributed w'ithin the tropical and subtropical zones (cf. Radwans!ki 1977a). A general conclusion is consequently drawn that the climatic con ditions prevailing within the Kortynica basin were identical with those typical at present or the tropical and/or subtropical zones. Within the other facies of the Fore-Carpathian Depression, especially within the detrital red-algal/bryozoan facies (Leithakalk type of the V1enna Basin), similar climatic requirements are indicated by the large echinoids Clypeaster, as well as by fish. Of the latter, both teleostean genus Scorpaena was reported (Jerzmailska 1958), as well as various elasmo branchs (Pawlowska 1960, Radwailski 1965b, Schultz 1977), of which the sharks Hemipristis serra Agassiz, and rays Aetobatis arcuatus Agassiz are the most tropical (Radwanski 1965b). All these fish, and the tropical elasmobranchs especially, may however by regarded as not the best climatic indicators, as they could migrate from warmer zones when being adult. The tropical andlor subtropical conditions in the Miocene seas of . Europe have previously been recognized by very few authors. Such conditions were concluded either fr9m the ecological analysis of parti cular genera, or from the whole organic assemblage contained in the deposits studied. To the first group belong conclusions drawn on the echinoid genus Clypeaster :investigated by Kalabis (1949) and Mltrovic ·Petrovic (1970), as well as on the gastropod genus Conus studied by Hall (1964) and Davoll (1972). The second type of analysis resulted. in the conclusions presented by Robba (1970) for the type Tortonian in Piedmont in Italy, and by Bohn-Havas (1973) for the Mecsek Mountains in southern Hungary. The Fore-Carpathian Depression is of all these regions situated undoubtedly the most northwardly (cf. Text-fig. lA), but lits. organic communities, these from the Korytnica basin including, do not differ in their composition from those indicated froOm southern Europe 4. It may therefore be postUlated that no evident climatic zonation 4 The only exception may be ascribed to the large gastropod species Pereiraia gervaisi (Yezian), which is. reported from south European countries (cf. Hoernes & Auinger 1879); its distribution of such a kind may however result from facies conditions (cf. Radwanski 1-975, p. 399). . 116 WACf..,AW BAf..,UK & ANDRZEJ RADWANSKI (cf. Radwanski 1975) was recognizable in the Miocene sea of Central Europe, that is, in the countries ranging from the southern Mediterranean as far as the Fore-Carpathian Depression. The problem, whether the - discussed tropical and/or subtropical conditions were prevailing only within the Miocene sea of Europe (due to, let us say, a pattern of warm currents), or they were stabilized over the Mioeene land, cannot be definitely solved yet. The recent biological and eoological investigations of the Miocene terrestrial plants of Europe indicate, in any case, that tropical elements were well pros pering within the then established floral communities. This is the best ciemOtl1Strated (Esike Koch & Friedrich 1971) by the zingiberaJcean species, Spiremato-spermum wetzleri (Heer) Chandler, the present-day :rIe1.atlive of which, Cenolophon oxymitrum (Schumann) Holttum, thrives oflly in relic sites in the jungles of Thailand. Similar climatic conditions were certalinly requdlred by tlhe gibbolns PUopithecu,.s commonlY' referenced from Central Europe -(cf. Zapfe 1960, Kowalski & Zapfe 1974). Both this banana-like plant, and the discussed ape are reported from Poland (cf. Eske Koch & Friedrich 1971; and Kowalski & Zap:£e 1974, respectively), the same as tropical/subtropical diving beetles of the genus Canthydrus recently found in siliceous sinter at Przeworno, Sudetlc Foreland (Galewski & Glazek 1973, 1977). The climate of the Polish land outside the Fa.re-Carpathian Dep:r:.ession (cf. Text-:Ilig. lA-B) should therefore be regarded as not deviated from that establiShed then in Central Eur,ope. The Korytnica basin with its very warm water; may be said to have been well suited within the tropi'C-influenced landscape of Central Europe. - I BIOGEOGRAPHIC APPLICATION Many of the tropical and/or subtropical elements that occur both in the Korytnica basin and in the other Miocene localities of southern and Central Europe '(Tethys and Paratethys basins: cf. Text-fig. lA), have Indo-Pacific affinities. Upon this statement, it was conduded (Radwanski 1974b, 1975) that from this very region the thermophilic elements penetrated to the Miocene sea of Europe. Such a route of their spreading is also evident from the composition of animal commu ruitdes in tJhe Miocene of Asia MinO!r, e.g., of the Sivas !region in eastern Turkey (cf. Stohepinsky 1939), and the Mut region in southern Turkey (cf. Sez.er 1970), both of which yield more tropical, Indo-Pacific elements that any of the above dis'Cussed localities in Europe. The SE:away connections through Turkey and the Persian Gulf regions are con sequently postulated, and these were in function until the crustaI up heavals resulted fntileir damming at the Miocene declli'ne (Radwatisld 1975; cf. also Baluk & RadwaIlski 1977). ORGANIC COMMUNITIES AND FACIES DEVELOPMENT 117 The Indo-Pacific affinities within the' organic assemblages of Central and Southern Europe are recognizable when comparing the present-day distribution of various genera, or of species which has been extant, and when analysing the whole communities. The first is . exemplified by such Cihitons as Cryptoplax (see references in Baluk 1971), many gastropods, and even the foraminifer Amphistegina lessonii Which the most widely distributed is in the Indo-Pacific (see referen.c€S in Rosenkrands Larsen 1976). The second is demonstrated by some of the seagrass communities (see references in Ffoffman 1977; cf. also Taylor 1971), and by an oyster-bryozoan-echinoicl-starfish community, reported from the locality Swiniary in the Fore-Carpathian Depression (Radwafislci 1970, 1973, 1974a, 1975), and well comparable to that known from the Gulf of MalllIlaT off north-western Ceylon (Herdrrum 1906). The Miocene North Sea Basin which belonged to the Atlantic province (cf. Text-fig. lA) bears also faunal influences of waters much warmer than at present (cf. Sorgenfrei 1958, pp. 41~15; Banke Rasmussen 1966, pp. 331-339; RadwaIlski & al. 1975; pp. 252-253). These influences are certainly indicative of water inputs from the tropical or subtropical part of the Miocene Atlantic Ocean (Gripp 1961, Radwanski 1975, Baluk & Radwanski 1977). These inputs were also responSible for the spreading of some thermophilic species in the Atlan tic gulfs of western. Europe (cf. Text-fig. lA), and few of suCh species could penetrate into the Te1lhys and Paratethys basins, as it is exempli fied by the cupufud-Jid bryozoons living in the KOil"'ytbIiIca baSin (cf. Cook 1965, Baluk & Radwanski 1977). These species should consequently be regarded (cf. Baluk & Radwailski 1977) as elements typical of the Atlantic province, although they lived tin the sea basins so influenced by even warmer waters of the Miocene ·Indo-Pacific. Institute of Geology of the Warsaw University, · Al. Zwirki i Wigury 93, 02-089 Warszawa, Poland REFERENCES BALUK, W. ill9l1.1. Lower Tortonian chitons from the Korytnica Clays, southern slopes of the Holy Cross Mts. Acta Geol. Polon., 21 (3), ~2. Warszawa. !9?2. Lower Tortonian scaphopods from the Korytnica Clays, southern slopes of the Holy Cross Mts. Acta Geol. Polon., 22 (3), 545-571. Warszawa. 1,9'75. Lower Tortonian gastropods from Kotytnica, Poland. Part 1. Pa,laeontol. Polon~, 32, 1-100. Warszawa-Krak6w. ,1977. 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